The role of birds and mammals in Korean pine (Pinus koraiensis) regeneration dynamics

This study examined the interrelationships of the fall seed-foraging guild with Korean pine (Pinus koraiensis) and its regeneration. The study took place in old-growth and secondary-growth forests in Northeastern China. Foraging behavior, seed transportation distance, and cache location of various birds and mammals were observed. Regeneration characteristics of Korean pine were also analyzed using plots in various vegetation cover types, successional stages, and topographical situations. Ten species of vertebrates were able to harvest seed from the closed-cone Korean pine. Of these, the Eurasian nutcracker, Eurasian nuthatch, red squirrel, and Siberian chipmunk were found to be potential seed dispersal agents. The nutcracker was the most important dispersal agent, easily acquiring seed with its large pointed bill, carrying up to 62 seeds in one trip, carrying seed at least 4 km, and placing seed in a variety of sites 2.5–3 cm deep in the soil. The Eurasian nuthatch carried a single seed per trip at distances less than 50 m, needed nutcrackers to open the cones and expose seed before they could acquire seed, and occasionally cached seed in the soil. Red squirrels were uncommon visitors to the tree tops of Korean pine, carried cones shorter distances than nutcrackers, and were only found under forest canopies. Human harvest of cones by knocking off branches also affected squirrel behavior and reduced future cone crops. Siberian chipmunks also collected seed from cones in trees and appeared to transport seed less than 50 m. Six other species were observed in this study successfully harvesting seed from cones but were not potential seed dispersers. Natural seedling establishment was found to be over 1000 seedlings/ha except on old-growth pine-hardwood sites. Squirrels were commonest here, but few seedlings survived past the 2nd year due to the intense shading. Second-growth forest types, including aPicea plantation where nutcrackers cached seed daily, and an old-growth pine-hardwood selective-harvest site, had the greatest regeneration. In conclusion, most natural regeneration of Korean pine in this part of its range is due primarily to the Eurasian nutcracker. Nutcrackers can aid forest managers in reaching desired stocking levels after disturbance, as well as a more natural-appearing forest. Squirrels, chipmunks, and nuthatches are minor seedling establishment agents. Korean pine seed is an important food source used by at least 22 species of forest wildlife.     

Responses in the physiology and biochemistry of Korean pine (Pinus koraiensis) under supplementary UV-B radiation

The effect of supplementary UV-B radiation on Korean pine (Pinus koraiensis Sieb. et Zucc) was investigated. Compared with the control, the T1, T2, and T3 UV-B treatments increased by 1.40, 2.81, and 4.22 kJ m^?2 d^?1, respectively. Gas-exchange parameters, photosynthetic pigment concentrations, contents of secondary metabolites, epicuticular wax, free radical, malondialdehyde (MDA), and the activities of antioxidant enzymes were determined after 40 d of exposure. The concentrations of chlorophyll (Chl) a, Chl b, total Chl, carotenoid (Car), and the ratio Chl a/b in the pine needles were in the following order: T1 > T2 > T3. Compared with the control, the contents of flavonoids and epicuticular wax significantly decreased in all levels of supplementary UV-B radiations (p<0.05). Moreover, the contents of hydrogen peroxide (H₂O₂) and MDA significantly increased with the enhanced UV-B radiations (p<0.05). Korean pine can increase the catalase, ascorbate peroxidase, and superoxide dismutase activities to prevent oxidative stress by supplementary UV-B radiation. However, its defence mechanism is not efficient enough to prevent UV-Binduced damage.     

Phylogeography of the Korean pine (Pinus koraiensis) in northeast Asia: inferences from organelle gene sequences

Range-wide genetic variation of Korean pine (Pinus koraiensis) was assessed using maternally inherited mtDNA and paternally inherited cpDNA for 16 natural populations throughout northeast Asia in order to study its phylogeographical history during the Quaternary. The cpDNA variation indicated that there was no difference between populations on the Asian continent and those in the Japanese archipelago. In contrast, the mtDNA variation indicated that there was significant difference between the populations from the two regions, with each region having a different lineage. The continental populations exhibited no diversity in the mtDNA examined despite the species’ current extensive range and large populations. Conversely, while the Korean pine is rare in Japan, the Japanese populations exhibited greater levels of mtDNA diversity (H _T?=?0.502). The higher mtDNA diversity and evidence from numerous Korean pine macrofossil remains dated to the Pleistocene and recovered various sites in Japan suggest that the Japanese archipelago once served as a refugium to a much larger Korean pine population with a more extensive range than is the case today. The presence of the single mtDNA haplotype across the Asian continent suggests that the present widespread populations could have expanded from a single refugium population after the last glacial periods.     

Fine root and litterfall dynamics of three Korean pine (Pinus koraiensis) forests along an altitudinal gradient

Background and aims

Fine root and aboveground litterfall, two large fluxes of nutrients and carbon in the forest ecosystems, are key processes to be considered in efforts of measuring, modeling and predicting soil carbon sequestration.

Methods

We used sequential coring and litter trap to measure seasonal dynamics of fine root and litterfall in three Korean pine dominated forests along an altitudinal gradient in the Changbai Mountain during the 2012 growing season.

Results

Fine root biomass decreased significantly while necromass increased remarkably with altitude. Patterns and amounts of fine root production and mortality varied among forest types. Litterfall decreased significantly with altitude, whereas forest floor mass increased. Carbon inputs through fine root mortality and litterfall decreased significantly with altitude while carbon storage of fine root mass did not differ among forest types and carbon storage of forest floor mass was significantly larger in higher altitudinal forests due to lower turnover rates.

Conclusions

This study provided an insight into the variations of fine root and litterfall dynamics among three Korean pine forests which were associated with different vegetation traits and environmental conditions, and also the quantification of carbon fluxes through fine root mortality and litterfall for estimating carbon budget of temperate forest.     

长白山阔叶红松林树木短期死亡动态

树木死亡是森林群落动态的重要过程,是多种因素共同作用的结果.本文基于长白山阔叶红松林25 ha样地2004年和2009年两次调查数据,从物种组成、数量特征、径级结构和空间分布等方面分析了5年间样地中死亡树木的特征.结果表明:5年间样地DBH≥1 cm的独立个体树种数由52变为51,3个树种因权有的1个个体死亡而消失,新增2个树种;独立个体数从36,908变为34,926,死亡个体数为4,030,死亡个体数占2004年个体总数的10.9%,新增个体数为2,048,独立个体数净减少1,982;死亡量大的树种其新增量也较大,灌木树种的死亡量和新增量均多于乔木树种;有5个树种的平均胸径减小,44个树种平均胸径增加;从死亡个体的径级结构来看,小径级个体死亡较多,大径级个体死亡少,5 cm以下的死亡个体占总死亡量的81.5%,不同林层的优势树种死亡个体的径级分布与2004年该树种的径级分布基本相同;不同树种死亡个体的空间分布具有较大差异,不同林层的优势树种死亡个体空间格局主要以聚集分布为主,小径级死亡个体在小尺度呈聚集分布,在其他尺度呈随机分布,中径级和大径级死亡个体在各尺度上都呈不同程度的随机分布.     

气候变化对阔叶红松林不同演替阶段红松种群生产力的影响

本文以长白山地区阔叶红松林不同演替阶段(次生杨桦林、次生针阔混交林、原始红松林)内红松种群作为研究对象,采用树轮学与相对生长式相结合,获取红松种群净初级生产力(NPP)连年生长(1921—2006年)数据以及相对增长率的年际变化数据,建立红松种群NPP与年际和季节性气候因子的关系,分析不同气候时期长白山阔叶红松林不同演替阶段内红松种群NPP年际变化特征及其对气候变化的响应差异.结果表明： 研究期间,不同演替阶段红松种群NPP与气候因子响应关系存在差异.随着温度上升,次生杨桦林红松种群NPP与上年生长季和当年生长季低温由显著负相关关系转变为显著正相关关系;次生针阔混交林红松种群NPP由与当年春季最低温度的正相关关系转变为与上年和当年生长季温度的显著正相关关系,气候因素对次生针阔混交林红松种群NPP影响的滞后效应增强;原始红松林红松种群NPP与温度的相关性减弱,与上年生长季降水量的正相关关系增强.研究区气候变化表现为低温和平均温度显著上升,而最高温度和降水没有明显变化.气候变化有利于提高演替初级阶段次生杨桦林和演替中级阶段次生针阔混交林内红松种群生产力,尤其是次生针阔混交林,而对演替顶极阶段红松种群NPP影响不明显.     

长白山阔叶红松林乔木树种幼苗组成及其年际动态

为了解阔叶红松(Pinus koraiensis)林乔木树种幼苗的组成及其年际动态, 以长白山阔叶红松林25 ha动态监测样地为平台, 在样地内150个种子收集器周围设置了600个5 m×5 m幼苗样方。基于2006–2008年连续3年的幼苗样方调查数据, 对乔木幼苗的树种组成、数量组成、空间分布特征、年际动态、新增和死亡幼苗组成等进行了分析。结果表明: (1)从树种组成来看, 该群落乔木树种的幼苗组成种类较为丰富, 共记录到21个树种, 这些树种也是样地内胸径1 cm以上乔木树种的主要组成成分。树种组成在年际间变化不大, 但各样方间表现出极大的空间变异。(2)从数量组成来看, 共记录到11,959株乔木幼苗, 以水曲柳(Fraxinus mandshurica)和紫椴(Tilia amurensis)幼苗数最多, 占总幼苗数的72.75%; 水曲柳、紫椴和红松的幼苗数量在年际间有明显波动, 其他树种年际间波动较小。(3)从新增和死亡幼苗的数量与组成来看, 共记录到15个乔木树种的新增幼苗, 其中紫椴、水曲柳、色木槭(Acer mono)、红松等10个树种在每次调查中都有新苗记录, 新苗数量在年际间随物种和样方位置表现出明显差异。(4)对各树种的幼苗、种子和大树的组成和空间分布的比较发现, 各树种的幼苗、种子和大树之间的数量组成和比例差异较大, 其中紫椴、水曲柳、色木槭和假色槭(A. pseudo-sieboldianum)的幼苗、种子在整个样地内都有分布, 春榆(Ulmus japonica)和怀槐(Maackia amurensis)幼苗的空间分布与种子和大树不一致, 糠椴(T. mandshurica)和山丁子(Malus baccata)等的幼苗、种子和大树的个体数相对都较少, 且它们的分布是一致的。     

长白山阔叶红松(Pinus koraiensis)林种子雨组成及其季节动态

为了解阔叶红松(Pinus koraiensis)林的种子雨组成及其在时间和空间上的动态变化,在长白山阔叶红松林25 hm2样地内,设置了150个种子收集器,收集掉落于种子收集器中的果实、种子等.所有收集到的样品分别鉴定其种类并分为成熟种实、未成熟种实、花序和果实或种子碎片及其附属物4类,计算各类别的数量,而后分别烘干秤重.从2005年6月到2006年11月,共收集到隶属11科12属20种的种实及其生殖器官残体.累计收集到121291粒种实,其中成熟种实23147粒,仅占所有种实总个体数的19.1%.种实数最多的树种是紫椴(Tilia amurensis)和水曲柳(Fraxinus mandshurica),两个树种种实的个体数占总个体数的90%.对2006年5月～11月种子雨季节动态的分析发现:种实在7月中旬至10月下旬数量极大,但主要由未成熟种实组成;在10月中旬出现成熟种实散落高峰,但未成熟种实仍占一定的比例.按每个收集器收集到的成熟种实数统计,成熟种实数量在100～200之间的收集器数量最多.按每个收集器收集到的树种数统计,收集器中最多收到的树种数为7,树种数为3和4的收集器个数最多.对6个主要树种成熟种实所在收集器的空间分布进行分析发现,紫椴和水曲柳的成熟种实在整个样地都有分布,春榆(Ulmus japonica)、糠椴(T. mandshurica)、色木槭(Acer mono)和假色槭(A. pseudo-sieboldianum)则只在样地的局部区域收集到成熟种实.成熟种实的空间分布与母树的空间分布大都表现出明显的相关性,表明这些树种的成熟种实并没有扩散到离母树较远的距离.     

Dynamic and allocation of recently assimilated carbon in Korean pine (Pinus koraiensis) and birch (Betula platyphylla) in a temperate forest

Biogeochemistry - Understanding how assimilated carbon (C) is allocated in forest ecosystems is crucial for process-based models and accurate prediction of global C cycling. We tracked the fate of...     

Different responses of Korean pine (Pinus koraiensis) and Mongolia oak (Quercus mongolica) growth to recent climate warming in northeast China

Different tree species growing in the same area may have different, or even contrasting growth responses to climate change. Korean pine (Pinus koraiensis) and Mongolia oak (Quercus mongolica) are two crucial tree species in temperate forest ecosystems. Six tree-ring chronologies for Korean pine and Mongolia oak were developed by using the zero-signal method to explore their growth response to the recent climate warming in northeast China. Results showed that Mongolia oak radial growth was mainly limited by precipitation in the growing season, while Korean pine growth depended on temperature condition, especially monthly minimum temperature. With the latitude decrease, the relationships between Korean pine growth and monthly precipitation changed from negative to positive correlation, while the positive correlation with monthly temperature gradually weakened. In the contrary, Mongolia oak growth at the three sampling sites was significantly and positively correlated with precipitation in the growing season, while it was negatively correlated with temperature and this relationship decreased with the latitude decrease. The radial growth of Korean pine at different sites showed a clearly discrepant responses to the recent warming since 1980. Korean pine growth in the north site increased with the temperature increase, decreased in the midwest site, and almost unchanged in the southeast site. Conversely, Mongolia oak growth was less affected by the recent climate warming. Our finding suggested that tree species trait and sites are both key factors that affect the response of tree growth to climate change. In addition, the suitable distribution area of Korean pine may be moved northward with the continued global warming in the future, but Mongolia oak may not shift in the same way.     

Composition and seasonal dynamics of seed rain in broad-leaved Korean pine (Pinus koraiensis) mixed forest in Changbai Mountain, China

To explore the composition and spatio-temporal dynamics of seed rain in broad-leaved Korean pine (Pinus koraiensis) mixed forest, 150 seed traps were set up in a 25 hm2 plot in Changbai Mountain. Seeds, fruits, anthotaxy and others in seed traps were collected, identified and divided into 4 types. From 2005 to 2006, we collected 47 different types. Total number of seeds and fruits was 121291, including 23147 mature seeds and fruits (19.1% of the total). Tilia amurensis and Fraxinus mandshurica, with the most seeds and fruits, accounted for 90% of the total. The analysis on seasonal dynamics of seed rain showed that there were the largest number of seeds and fruits between July and October, which were composed of immature seeds and fruits. In mid-October, mature seeds and fruits reached their peak, but immature seeds and fruits still accounted for high proportion. There were 91 traps that contained 100–200 mature seeds and fruits, and one trap without any mature seed or fruit. The largest number of species found in a trap was 7, and usually 3 or 4 species were found in most of the traps. There were obvious relationships between spatial patterns of mature seeds and fruits and their parent trees, indicating that their mature seeds and fruits were not dispersed far from their parent trees.     

长白山阔叶红松林的温度效应

森林温度特征的研究是揭示森林生态系统功能、评估森林环境效益的基础.对2004年长白山阔叶红松林林内外空气温度和土壤温度进行了观测,结果表明:林内外空气温度、土壤温度明显不同.白天林内气温一般低于林外,夜间高于林外.昼间气温差夏季较大,7月平均差值为1.6 ℃;夜间气温差冬季较大,1月平均差值为1.5℃.空气温度和土壤温度的日振幅总是林内小于林外,7月空气温度日振幅林内比林外低1.7℃,4月土壤温度日振幅林内比林外低5.8℃.     

长白山阔叶红松林物种多度和空间分布格局的关系

应用随机分布多度模型和聚集分布多度模型,探讨不同研究尺度下物种多度和空间分布格局的关系.结果表明,预测的物种多度不仅受物种分布面积大小的影响,还受其聚集程度的影响.物种多度和空间分布格局的关系存在着明显的尺度效应,即随着研究尺度的增加,无论是随机分布多度模型还是聚集分布多度模型,通过物种空间分布格局来预测物种多度的准确度都在下降.聚集分布多度模型预测物种多度的结果要好于随机分布多度模型,这表明该区大多数物种是聚集分布的.由于物种的空间分布格局不同,不同物种多度的预测值和真实值之间的差异也不同.因此,为了进一步提高模型预测的准确性,进一步考虑不同物种的生活史特性是必要的.     

长白山阔叶红松林草本植物多样性季节动态及空间分布格局

基于长白山阔叶红松林25 hm²样地(CBS)一年内草本植物的4次调查数据,对样地内草本植物多样性的季节动态及其空间分布格局进行了初步分析.结果表明：样地内草本植物物种组成丰富,共有102种,隶属于40科84属.Shannon多样性指数、Simpson多样性指数和Pielou均匀度指数分别为3.52、0.96和0.75;物种组成的季节变化比较明显,以初夏物种数最多;各季节的Shannon多样性指数、Simpson多样性指数和Pielou均匀度指数的变化较大,个体数量从早春到秋季逐渐减少;物种丰富度和多度的空间分布连续性较差,主要表现为斑块性分布,说明草本植物对微环境有较强的依赖性;坡向是影响物种丰富度和多度的主要因素, 在早春、夏末和秋季,不同坡向的物种丰富度和多度差异极显著(P<0.01),且早春阶段北坡和东坡的丰富度高于南坡和西坡,夏末和秋季则相反.     

长白山阔叶红松林灌木物种组成、结构和空间分布

以长白山阔叶红松林25 hm2动态监测样地为平台,基于样地内600个5 m×5 m样方的调查数据,对灌木的物种组成、基本特征、结构和空间分布等进行了初步研究.结果表明:共调查到18种灌木,以东北山梅花、毛榛和簇毛槭为主要优势种;调查样方共记录灌木个体6435个和灌木枝干11369个,其中东北山梅花的个体数和枝干数最多,分别占总数的40.6%和33.4%;不同灌木的成丛率、每丛枝干的数量、冠幅和基径均存在明显的差异;不同灌木在高度上有较为明显的分化,灌木群落成层现象明显;优势种东北山梅花分布广泛,而珍珠梅、石蚕叶绣线菊、柳叶绣线菊和瘤枝卫矛的空间分布则呈现出较明显的异质性.     

Photosynthesis, leaf morphology and chemistry of Pinus koraiensis and Quercus mongolica in broadleaved Korean pine mixed forest

Leaf traits and physiology are species-specific and various with canopy position and leaf age. Leaf photosynthesis, morphology and chemistry in the upper and lower canopy positions of Pinus koraiensis Sieb. et Zucc and Quercus mongolica Fisch. ex Turoz in broadleaved Korean pine forest were determined in September 2009. Canopy position did not significantly affect light-saturated photosynthetic rate based on unit area (P _area) and unit dry mass (P _mass), apparent quantum yield (α), light compensation point (LCP), light saturation point (LSP); total nitrogen (N_m), phosphorus (P_m), carbon (C_m), and chlorophyll content (Chl_m) per unit dry mass; leaf dry mass per unit area (LMA) and photosynthetic nitrogen-use efficiency (PNUE) for P. koraiensis current-year needles and Q. mongolica leaves. While in P. koraiensis one-year-old needles, P _area, P _mass, α and LCP in the upper canopy were lower than those in the lower canopy. The needles of P. koraiensis had higher Cm and LMA than leaves of Q. mongolica, but P _mass, Chl_m and PNUE showed opposite trend. There were no differences in P _area, LSP, N_m, and P_m between the two species. Needle age significantly influenced photosynthetic parameters, chemistry and LMA of P. koraiensis needles except LCP, LSP and C_m. In contrast to LMA, P _area, P _mass, N_m, P_m, Chl_m, and PNUE of one-year-old needles were significantly lower than those of current-year needles for P. koraiensis. The negative correlations between LMA and P _mass, N_m, P_m, Chl_m, and positive correlations between P _mass and N_m, P_m, Chl_m were found for P. koraiensis current-year needles and Q. mongolica leaves. Our results indicate that leaf nitrogen and phosphorus contents and nutrient absorption from soil are similar for mature P. koraiensis and Q. mongolica growing in the same environment, while difference in carbon content between P. koraiensis and Q. mongolica may be attributed to inherent growth characteristics.     

长白山阔叶红松林凋落物组成及其季节动态

为了解群落尺度上凋落物组成及其时空变化,对长白山阔叶红松林25 hm² 样地内2008年度收集的凋落物进行统计分析.结果表明: 一年间,150个收集器内收集的凋落叶分属35种树木,占样地内胸径≥1 cm树种数(52种)的67.3%;凋落物总量为29.39 kg,折合3918.4 kg·hm^-2,其中,阔叶、杂物、针叶和枝条凋落量分别占凋落物总量的61.7%、18.0%、11.7%和 8.6%;紫椴、水曲柳、蒙古栎、色木槭和春榆5个树种的叶凋落量占阔叶总凋落量的83.8%;不同树种的凋落量季节差异很大,61.9%的凋落物产生于9月13日至10月10日.其中,红松和紫椴叶凋落高峰出现在9月13-26日,蒙古栎、春榆和色木槭叶凋落高峰出现在9月27日-10月10日.收集器间凋落物量差异较大,其中68个收集器的年凋落量在150～200 g,1个收集器大于500 g;单个收集器全年最多可收集到18个树种的凋落叶,凋落叶种数为12种的收集器最多(32个).叶凋落量与样地内该树种的胸高断面积总和成正比.样地内凋落叶的分布存在明显的空间异质性,且收集器内凋落物的收集量与其所处的位置有关.     

Newly-formed photosynthates and the respiration rate of girdled stems of Korean pine (Pinus koraiensis Sieb. et Zucc.)

A stem-girdling experiment was carried out on an evergreen conifer, the Korean pine (Pinus koraiensis Sieb. et Zucc.), in mid summer in Northeast China. A 50 % higher respiration rate at the upper part of the stem was observed 3 d after stem girdling, and a stable higher rate (1.2–2.8 times) one week later. However, no higher soluble sugar or starch contents were found in the upper bark of the girdled stems in measurements over three weeks. These findings indicate that most of the newly-formed photosynthates were consumed by the high respiratory activity; this is also implied by the strong correlation between the photosynthetic photon flux over the canopy (PPF) and respiration at the upper parts of girdled stems. Moreover, the maximum PPF and cumulative PPF one day before measurement (PPFmax-Y and CPPF-Y, respectively) were closely correlated with the respiratory difference between the upper and the lower parts, but no such correlation was found with the instantaneous PPF (PPF-I) and cumulative PPF on the current day from sunrise to measured time point (CPPF-C). This shows that photosynthates newly formed by canopy needles need at least one day for transportation in order to increase the stem respiration at tree breast height.     

长白山阔叶红松林红松种群年龄结构与数量动态特征

该研究以分布于长白山阔叶红松林内的红松(Pinus koraiensis)种群为对象, 通过编制种群静态生命表, 计算数量动态指数, 绘制存活曲线、死亡率曲线和消失率曲线, 应用4个生存函数并引入谱分析和时间序列预测模型, 分析红松种群年龄结构, 揭示其天然更新过程及未来发展趋势, 以期为野生红松种群的保护和恢复提供科学依据。结果显示: 红松种群数量变化具有阶段性, 幼龄(I-III龄级)和成龄(VII-X龄级)个体数量多, 中龄(IV-VI龄级)和老龄(XI-XIV龄级)个体数量少, 形成明显间断的两个优势年龄分布区。种群存活曲线趋近于Deevey-Ⅲ型, 表明幼龄个体死亡率高。忽略外部干扰时的总体数量动态变化指数大于>0, 表明红松种群为增长型; 考虑未来外部干扰时的种群动态变化指数趋近于0, 结合死亡率和消失率曲线呈现出连续先增后降的复杂动态变化趋势, 可知该种群受外界随机干扰时增长不明显。生存函数分析显示, 红松种群具有前期锐减、中期稳定、后期衰退的特点。谱分析表明红松种群天然更新呈周期性波动。未来2、3、4、5、6、7、8、9、10个龄级时间后, 红松幼、中龄个体数逐渐减少, 而成、老龄个体数量将逐渐增加。幼龄个体死亡率高、生存空间和资源条件有限, 老龄个体生理衰老明显是限制红松种群增长的主要原因。建议加强幼龄个体的抚育工作, 提高其存活率和生存质量; 保护和改善生存环境, 从而促进红松种群的自然更新和恢复。     

Seed dispersal of the Korean pine,Pinus koraiensis,by the red squirrel,Sciurus vulgaris

Seed hoarding behavior of the red squirrel,Sciurus vulgaris, was studied in relation to the amount of dispersed seeds of the Korean pine,Pinus koraiensis, and the distribution of its seedlings. After removing a cone from a tree, squirrels sat on the ground and ripped off its cone scales before transporting it. A mean of 3.2 seeds were scatter-hoarded per hole. Of 7.7×10⁴ mature seeds produced in a 0.21 ha planted Korean pine forest, 22% were estimated to be directly eaten by four squirrels, 9% were hoarded by them in the pine forest and 65% were cached outside the forest. Squirrels rediscovered hoarded seeds frequently, until the ground was covered with snow, during the period from snow fall until seed germination the next spring, few hoarded seeds were utilized. Korean pine seedlings were found up to 600 m from their mother trees. Scatter-hoarding by squirrels extensively contributes to seed dispersal to places suitable for the regeneration of the Korean pine. The large size of the cone, the absciss-layer at the cone penduncle, the infrequent dehiscence of cone scales, the large and wingless seeds, and the thick seed-coats have probably all been specialized to facilitate utilization by GenusSciurus.